Scientific Research and Essay Vol.4 (10), pp.

997-1005, October, 2009

Available online at http://www.academicjournals.org/sre
ISSN 1992-2248 © 2009 Academic Journals

Full Length Research Paper

Comparison of Streptomyces diversity between

agricultural and non-agricultural soils by using various
culture media
Mustafa Oskay
Biology Department, Faculty of Sciences and Arts, Celal Bayar University, Manisa, Turkey. E-mail:
mustafa.oskay@bayar.edu.tr
Accepted 18 August, 2009

Streptomycetes play a key role in the sustainability of agriculture and indicates the level of health of
soil, especially when considering the richness of them that are involved in biological control of soil
borne diseases. 20 different soil samples were taken from agricultural (7) and non-agricultural places
(13) and populations of streptomycetes were quantified in order to select the general culture media that
had better reflect the changes of these bacteria. The most efficient medium for the isolation of
Streptomyces was starch casein agar by the addition of nystatin. Pretreatment of soil samples with
CaCO3 (1%) increased the streptomycetes occurring on the isolation plates. To establish a correlation
with soil physico-chemical parameters, such as pH, salt, N, P, K, Na, Fe, Zn and Cu were also
determined, most of the correlations being significantly positive on the quantification of Streptomyces
6 6
diversity. Streptomycete counts ranged from a high of 6.7 x 10 to a low of 2.3 x 10 cfu/g dry soil of
non-agricultural soils. Streptomycetes constituted 4.8 to 45.8% of the total culturable bacterial
community. Higher streptomycete densities were greatest in non-agricultural soils with an average of
14.0% compared to agricultural soils with average of 10.1%. These results suggest that these bacteria
may be represent an unexplored resource for pharmaceutical drug discovery but also may provide
additional disease control in agriculture.

Key words: Agricultural soil, non-agricultural soils, culturable bacteria, isolation, pretreatment, Streptomyces
diversity.

INTRODUCTION

The soil microbes perform a wide range of function in the
ecosystem. Among soil organisms, bacteria and fungi,
actively participate in organic matter decomposition
liberating chemical nutrients and furthering plant growth.
Bacteria and fungi also play an important role for the
stability and productivity of agricultural soils. Therefore for
a sustainable and nature saving agriculture with high crop
yields the genetic diversity of microbes may play an
important role and could be used as an indicator for soil
quality, which has to be determined for understanding
turnover processes (Schloter et al., 2003; Vargas Gil et
al., 2009a). Therefore, soil microbes have an important
role to the subsistence on earth, because it has the role
on biological and chemical cycling among the flora, fauna
and life of microbes itself. Each type of microbes fills as a
unique niche, plays a different role in nutrients cycling
and soil structure. Microbial communities may be used as
indicators of the ecological equilibrium between pathogens
and biocontrol agents naturally suppress the incidence of
diseases. Since microbial diversity includes the number
of different fungal and bacterial species and their relative
abundance (Lartey, 2006; Martinez Blanco et al., 2007;
Vargas Gil et al., 2009b). Growth of microbial populations
and their action on soils are dependent on the interaction
between soil type, plant species and its rhizosphere
localization. Microorganism numbers vary in and between
different soil types and conditions, with bacteria being the
most numerous (Vieira and Nahas, 2005).
The isolation and characterization of pure cultures of
some microbial species are as important as under-
standing their objective existence in natural ecosystems.
The isolation of diverse and novel pure cultures of
actinomycetes provides a theoretical guide for the exploit-
tation and utilization of actinomycete resources (Li et al.,
1996). Streptomyces are one of the groups of actinomy-
cetes that are widely spread in both terrestrial and aqua-
998 Sci. Res. Essays
tic environments (Cross, 1989; Locci, 1989; Williams et
al., 1989). They play an important role in the circulation of
organic substances in nature. Many representatives of
this group have important practical interest as producers
of antibiotics and other biologically active substances of
high commercial value and are being routinely screened
for new bioactive substances, which have wide use in
biotechnological production (Anderson and Wellington,
2001; Vijayakumar et al., 2007). Their distribution and
predominance depends mainly on several factors, such
as nutrient availability, temperature, pH, moisture, soil
type, season and climate (Waksman, 1961; Williams et
al., 1989; Katsifas et al., 1999; Saadoun and Gharaibeh,
2003). Streptomyces have been found in all known soils
of the world yet their number, role in biocenosis and
biochemical activity vary depending on ecological and
geographical conditions (Dolotkeldieva and Totubaeva,
2006). One of the aims of biodiversity studies of action-
mycetes is to use effective isolation procedures to study
the distribution of actinomycetes in various climatic and
ecological environments (Li et al., 1996). Many studies on
the ecological distribution of soil streptomycetes and their
biotechnological importance in detail has been reported
from normal and agricultural soils (Conn and Leci, 1998;
Katsifas et al., 1999; Mitra et al., 2008; Zenova et al.,
2008; Grishko and Syshchikova, 2009; Vargas Gil et al.,
2009b).
Some soil microorganisms have been studied in detail
however; studies that are more comprehensive are nee-
ded to understand the diversity, distribution and ecology
of the large majority of streptomycetes in terrestrial
habitats. The objectives of this study were to compare
and describe the diversity of Streptomyces from different
agricultural and non-agricultural fields, which have
different crops and vegetation in relation to soil chemical
properties. These areas are yet poorly studied and repre-
sent diverse and largely unscreened ecosystem for the
isolation of Streptomyces that potent for pharmaceutical
industry and agriculture.
MATERIALS AND METHODS
Soil sampling and processing
7 soil samples were collected from agricultural fields (AG) (Corn,
cotton, wheat, barley, vegetable, vineyard and orchard fields) in
Manisa Province, Turkey and 13 of non-agricultural soils samples
(NAG) were collected from various locations of North Cyprus and its
surroundings into sterile plastic bags, to avoid external conta-
mination.. Every sample is a mixture of soils from five to ten holes
at a depth of from 10 to 30 cm. Soils were air-dried and stored at
4.0°C until processed.
Calcium carbonate soil treatment
A method (El-Nakeeb and Lechevalier, 1963) with some modify-
cations was used according to our laboratory conditions. The air-
dried soil (10 g) was mixed in a mortar with 1% of calcium carbon-
carbonate (CaCO3) and then incubated for 2 days at 30.0°C in a
closed inverted sterile Petri dish in which a high relative humidity
was maintained by water saturated of filter paper. To assess the
effect of pretreatment, soils without CaCO3, served as a control.
Isolation and enumeration of Streptomyces
10 g of pretreated soil samples were added to 90 ml sterilized water
in 250 ml Erlenmeyer flasks. Flasks were shaken on rotary shaker
at 200 rpm for 30 min. All samples were diluted (up to 10-7) with
sterile distilled water prior to inoculation into the isolation plates.
Isolation of streptomycetes were performed by soil dilution plate
technique using different media such as starch-casein agar (SCA)
(Kuster and Williams, 1964), glycerol asparagine agar (ISP 5)
(Shirling and Gottlieb, 1966), potato dextrose agar (PDA) and
nutrient agar (NA). All isolation media also contained nystatin at
final concentrations of 50 µg/ml, to minimize fungal contamination
(Waksman, 1961). The pH of each medium was adjusted to 7.0 -
7.5 to match that of the soil sample. 1 ml of soil of 10-6 dilution (in
most situations) is plated out and thoroughly mixed with about 20 -
25 ml of melted desired agar medium at around 45.0 - 50.0°C. After
gently rotating, the isolation plates were incubated at 28.0 - 30.0°C
for 7 - 14 days to allow sufficient time for fast-growing
streptomycetes or longer for the slow-growing ones.
Streptomyces counts
Streptomycetes were quantified on each plate by eye and with the
aid of a stereomicroscope (Olympus, magnification: 10 - 90 x); then
recognized by the presence of filamentous hyphae; a characteristic
that was just within the range of detection at the highest magni-
fication used and/or by the formation of floccose, powdery, tough,
leathery colonies that adhered to the agar surface and colors of
pigmentation including diffusible pigments (Waksman, 1961;
Williams et al., 1989; Cross, 1989; Anderson and Wellington, 2001).
Colony formation units (cfu) per gram counting the average for each
soil sample estimated for densities of total culturable streptomy-
cetes and bacteria. The total number of streptomycete colonies
observed was counted and representatives with different morpho-
logies were obtained in pure culture by repeated transfer from a
single colony. Slants of yeast extract-malt extract agar (ISP2) or
oatmeal agar (ISP4) containing pure cultures were maintained at
4°C in culture collection of Biology Department, Celal Bayar
University, Manisa Turkey.
Physico-chemical analyses of the soil samples
Samples were also taken from each site for analyzing physico-
chemical parameters such as soil structure, lime (CaCO3), satura-
tion, pH, salinity, available nitrogen (N), phosphorus (P), potassium
(K), sodium (Na), ferrous (Fe), copper (Cu), zinc (Zn), manganese
(Mn), calcium (Ca) and magnesium (Mg) (Scheffer and
Schachtschabel, 1966; Schlichting and Blume, 1966; Ryan et al.,
1996). The air-dried soil samples were ground mixed properly and
sieved to remove gravel and debris. Physico-chemical parameters
of soils were determined in the Vali Ecemi Soil Analysis Laboratory
of Manisa, Directorate of the Ministry of Agriculture (report number:
3644/63-06; report date: 20.12.2006).
Taxonomic grouping of isolates
Streptomyces colonies were placed in genera and taxonomic
groups based on the morphological, cultural characteristics and
chemical compositions of cells. Morphological and cultural observa-
tion were carried out by using the methods and media proposed as
described in the International Streptomyces Project (ISP) (Shirling

and Gottlieb, 1966) and the Bergey’s Manual of Systematic Bacte-
riology (Cross, 1989; Williams et al., 1989). The morphology of the
spore bearing hyphae with the entire spore chain, the structure and
arrangement of the spore chain with aerial mycelium of the
streptomycetes were examined using slide culture technique
(Cross, 1989). After growth, the slide cultures were examined under
light microscope (Magnification, 400 and 1000X). Colors were
determined according to the scale adopted by Prauser (1964) and
isolates were grouped into separate color series according to the
system proposed by Shirling and Gottlieb (1966). For chemotaxo-
nomic studies, aerial and substrate mycelia of streptomycetes were
scraped from the ISP2 plates and processed for the isomers of
diaminopimelic acids (LL-DAP or meso-DAP) and whole cell sugar
patterns by the method of Lechevalier and Lechevalier (1970).
Precoated silica gel plates (20X20, 60 F , Merck, Darmstadt,
254
Germany) were used for thin layer chromatography.
Statistical analysis
The average values of the number of cfu g/dry soil were statistically
analyzed by Minitab 13.20 (Minitab Inc., 2000) program by the
multivariate cluster analysis to find out the similarity (%) of Strepto-
myces diversity between agricultural and non-agricultural soils.
RESULTS AND DISCUSSION
Streptomycetes are best known as soil bacteria and
largely occur as dormant spores (Waksman, 1961; Cross,
1989; Williams et al., 1989). The distributions and
ecological roles of Streptomycetes in the agricultural
environment have remained an unresolved issue in soil
biology. In an effort to gain a better understanding of soil
streptomycete diversity, a culture-dependant study was
undertaken using samples collected from agricultural and
non-agricultural soils.
The occurrence of streptomycetes in the 20 soil sam-
ples (13 from non-agricultural and 7 from agricultural soil)
were extensively investigated using CaCO3 treatment
procedure and different media. The distributions of
colonies in four different media were expressed by sum-
−6
ming up the colonies observed of the 10 dilution series.
6 6
Mean of Streptomyces colonies were 4.1 x 10 (14.0%)
6
and 3.3 x 10 (10.1%) for NAG and AG, respectively.
However, the total isolated bacterial colonies (adding all
the colonies obtained in four different media) were 35.5 x
6 6
10 and 29.5 x 10 for NAG and AG, respectively (Table
1). The influence of culture media on streptomycete
diversity was evaluated. Among the four different media
used, SCA was the most effective medium supplemented
with nystatin (50 µg/ml) in the isolation of streptomycetes
6
from soils (average, 5.4 x 10 cfu/g). PDA was the se-
6
cond most effective (3.7 x 10 cfu/g) while GAA, was the
6
third (3.3 x 10 cfu/g). NA was the least effective medium
6
for the isolation of streptomycetes (2.6 x 10 cfu/g)
(Figures 1 and 2). Varied nature of growth of the strepto-
mycetes, chemical parameters of soils (Table 2) and the
type of selective media used may be the three possible
reasons for this result. The amount of time for Strepto-
myces colony development was similar in all culture
media tested. The use of nystatin in culture media notably
Oskay 999
reduced fungal contamination; it was effective in the
count of streptomycetes. Total numbers of Streptomyces
in each soil samples of CaCO3 treatment were expressed
as the average number of colonies in four different media
(Table 1).
In the control experiment, a significantly lower amount
of streptomycetes was recorded, compared with CaCO3
treatment. Average of Streptomyces densities were
6 6
recorded 1.9 x 10 (5.0%) and 1.6 x 10 (3.9%) for NAG
and AG, respectively. However, the amount of total cul-
6 6
turable bacteria increased as 42.5 x 10 and 38.6 x 10
for NAG and AG, respectively.
Results revealed that the streptomycetes diversity of
the sampling sites was influenced by the chemical nature
of the soil. A very typical pattern of soil characteristics of
almost all of the samples were observed, that is low in
salinity, P and Zn (except the sample site, ST8) (Table 2).
Among the exchangeable cations (Fe, Na, K, P, Cu, Zn,
+
Mn, Ca and Mg), Ca (4256 ppm) was present in the
+ + + + +
highest amount following Na , Mg K , Fe and Mn . The
soil pH was ranging from neutral to slightly alkaline (pH
range 7.23 to 7.78). Soil of the ST8 contains higher
amounts of P, which supports the proliferation of Strepto-
myces (Parfitt et al., 2005). The total N was observed
between 1.33 ppm in ST3 and 134.43 ppm in ST14. The
highest of Streptomyces colonies were observed in site,
6
ST8 (6.7 x 10 cfu/g), having rich N, P, Fe, Mn and Ca;
clayed-loamy soil and the lowest number in ST15 (1.9 x
6
10 cfu/g), that site containing wheat vegetation. The
highest number of total viable bacteria was observed in
6 6
ST9 (48.1 x 10 cfu/g) and the lowest in ST18 (20.4 x 10
cfu/g) of vineyard soils.
There was significant variation in total viable bacterial
and Streptomyces densities between field types in vitro.
Overall, streptomycete densities were negatively corre-
lated with total bacterial densities and streptomycete
densities were positively correlated with vegetations.
Similar patterns were found within each soil type. Corn-
fields had a marginally significantly greater density of
streptomycetes (6.2 x 10 cfu/g) than other agricultural
fields that had streptomycete density ranging from 1.9 to
6
3.8 x 10 cfu/g. However, according to the all-statistical
analyses, there was not relationship between the
intensity of streptomycete in AG and NAG fields, which
were similar percentage at least 98.47% (Figure 3). This
result suggest that just a small percentage of soil
streptomycete is culturable (1 - 3%) (Watve et al., 2001),
even using a set of culture media (Waksman, 1961).
Therefore, using a selective media is a highly necessary
step, because Streptomyces densities are directly affect-
ted by culture medium. However, considering the diffe-
rent of culture media and many environmental factors,
most streptomycetes can use a wide variety of com-
pounds as energy source such as glucose, starch, amino
acids and proteins. The best nitrogen sources for these
bacteria are proteins, peptones, amino acids, nitrates and
ammonium salts (Waksman, 1961; Shirling and Gottlieb,
1966). In addition, the main factors that determine
1000 Sci. Res. Essays

Table 1. Comparison of culturable Streptomyces diversity and total bacteria between NAG and AG.

Calcium carbonate treatment No treatment

Sites Culturable Total viable Streptomyces Culturable Total viable Streptomyces
Streptomyces bacterial Streptomyces bacterial
counts counts % counts counts %
b a
ST1 5.2 28.6 18.1 2.7 36.7 7.4
ST2 4.7 32.8 14.3 2.2 34.4 6.4
ST3 4.4 36.4 12.0 1.4 48.6 2.9
ST4 3.3 38.7 8.5 2.2 44.3 5.0
ST5 2.6 40.2 6.5 1.3 41.2 3.1
ST6 3.5 41.3 8.5 1.6 46.6 3.4
ST7 4.8 30.4 15.8 2.6 33.9 7.7
c
ST8 6.7 22.3 45.8 3.8 36.8 10.3
ST9 2.3 48.1 4.8 0.9 58.4 1.5
ST10 4.2 32.6 12.9 1.3 40.5 3.2
ST11 2.7 39.5 6.8 1.1 45.2 2.4
ST12 2.3 42.2 5.4 0.8 53.7 1.5
ST13 6.4 28.3 22.6 3.4 33.4 10.2
ST14 6.2 36.2 17.1 2.8 48.9 5.7
ST15 1.9 24.6 7.7 0.9 30.1 3.0
ST16 2.2 33.7 6.5 1.1 33.8 3.2
ST17 3.8 31.3 12.1 2.6 43.7 6.0
ST18 1.8 20.4 8.8 0.7 28.6 2.4
ST19 3.6 29.2 12.3 1.7 40.3 4.2
ST20 3.3 30.6 10.8 1.3 44.8 2.9
a
Average of 40 plates of four different media, viable counts = x106 cfu/g dry soil.
b
Samples of ST1-ST13 are non-agricultural soils, ST14-ST20 are agricultural soils (ST14: Corn, ST15: cotton, ST16: wheat, ST17: barley,
ST18: vegetable ST19: vineyard and ST20: orchard fields).
c
highest values are indicated with bold, lowest values are given bold and italic.

Figure 1. Comparison of the influence of culture media on culturable Streptomyces counts (Average of the
total Streptomyces populations of AG and NAG soils; counts from 200 plates of each medium; cfu: colony-
forming units).
 Oskay 1001

Figure 2. Streptomycete colonies on four different media, SCA; starch casein agar (a-b), PDA; potato
dextrose agar (c-d), GAA; glycerol asparagine agar (e), NA; nutrient agar (f), in addition –UT; untreated, –T;
treated with CaCO3, dilution; 10-6. Note that the majority of colonies on these media are Streptomyces
bacteria that are easily recognized by their spherical and wrinkled shape, diffusible pigments and colonies
in different color (here white, grey, yellow, cream and pink or red; examples of Streptomycete colonies are
shown by green arrows). Photographs were taken after plates had been incubated for 2 weeks at 28.0°C.
 1002 Sci. Res. Essays

Table 2. Physico-chemical properties of agricultural and non-agricultural soil samples (10 - 30 cm).

Non-agricultural soils Agricultural soils

Soil property
ST1 ST2 ST3 ST4 ST5 ST6 ST7 ST8 ST9 ST10 ST11 ST12 ST13 ST14 ST15 ST16 ST17 ST18 ST19 ST20
a
Soil structure C-L C-L L C-L L L L C-L L L C-L C-L C-L C-L C-L C-L C-L C-L L L
Salinity (µS/cm)b 686 534 477 1103 199 675 2570c 1327 576 632 475 302 780 2300 723 541 308 268 319 238
Lime (%) 14.82 1.95 17.55 17.55 11.70 2.11 2.73 10.92 17.16 19.50 3.51 5.07 18.33 14.82 12.48 15.21 3.12 3.51 2.73 4.68
Saturation (ml) 58 59 50 51 44 50 50 56 49 49 51 58 55 69 68 56 53 49 42
pH 7.78 7.53 7.37 7.32 7.55 7.28 7.35 7.23 7.70 7.20 7.54 7.77 7.40 7.64 7.49 7.44 7.47 7.63 7.50 7.44
N (ppm) 1.38 1.52 1.33 1.80 1.62 2.23 1.65 16.85 1.73 2.14 2.12 3.27 12.26 134.43 2.22 2.68 2.05 1.54 2.06 1.59
P (ppm) 0.32 2.61 1.52 8.65 2.69 3.14 0.10 16.77 4.10 3.47 1.56 8.21 5.66 9.12 7.86 15.09 12.68 7.81 20.61 6.69
K (ppm) 263 176 226 263 205 389 169 298 2.0 246 186 165 297 433 359 375 342 214 287 181
Na (ppm) 310 119 115 160 42 166 554 527 33 147 117 119 230 120 81 57 27 45 28 19
Fe (ppm)d 5.31 3.47 3.91 4.30 5.07 4.35 4.67 53.80 60.20 5.19 4.43 9.35 5.35 3.71 5.03 7.02 2.51 6.67 8.66 4.30
Cu (ppm) 1.39 3.27 2.51 2.96 2.10 2.38 1.73 2.09 5.32 2.97 3.56 2.71 3.32 2.75 2.66 2.39 2.66 7.24 13.25 5.37
Zn (ppm) 0.14 0.42 0.24 0.89 0.21 0.57 0.12 2.66 0.82 0.52 0.71 2.81 0.75 0.89 0.69 1.95 0.82 0.70 1.36 3.73
Mn (ppm) 2.68 6.10 1.26 2.60 2.98 5.40 1.09 20.08 12.36 2.49 6.74 2.93 3.29 9.51 2.09 28.12 3.76 2.17 6.75 2.77
Ca (ppm) 4200 3200 3288 4256 2894 2866 3267 4020 2967 3111 2914 3098 4024 3867 3714 3024 2916 3214 2866 3024
Mg (ppm) 364 244 248 428 255 266 312 420 291 314 290 302 288 266 388 294 301 299 294 304
a
Soil structure, C-L: Clayed-loamy soil, L: Loamy soil.
b
µS/cm: Microsiemens per centimeter, ppm: Parts per million.
c
First three highest values are indicated with bold for parameters of each site.
d
Fe, Mn, Zn and Cu: extracted with ammonium bicarbonate-diethylenetriamine pentaacetic acid (AB-DTPA).

streptomycetes development are pH and tempera-
ture (Williams et al., 1983).
Soil dilution and isolation on culture media have
proved to be a useful method; however, it has
some limitations: the methodology is slow and
laborious and requires a large volume of material.
Hence, given the small cultu-rable portion of soil
microorganisms, any biodiversity study is limited;
furthermore, the nutritional and physio-logical
requirements of streptomycetes can be so specific
that they restrict the study even more (Waksman,
1999;
Williams et al., 1989; Vargas Gil et al., 2009b).
According to morphological and cultural obser-
vations indicated, that taxonomic grouping of
these isolates is in the genus Steptomyces as re-
ported by other researchers (Shirling and Gottlieb,
1966; Williams et al., 1983; Anderson and
Wellington, 2001). The compositions of DAP and
sugar components of selected representative’s
isolates were detected. DAP existed as isomers
with LL types. Cells contained no diagnostic sugar
components. Streptomyces isolates were catego-
rized into six color series according to the color of
their mature sporulated aerial mycelium with grey
and white color series being the most abundant
(Figure 4). Data indicated that 38 and 25% of
NAG; 30 and 32% of AG fields have grey and
white color series of Streptomyces, respectively.
Saadoun and Gharaibeh (2003) and Thakur et al.
(2007) reported similar results; they showed that
grey and white series of Streptomyces had the
highest occurrence in soils. Microscopic examina-
tion of the spore morphology revealed that most of
the isolates had rectiflexibile spore type (55 and
45% for NAG and AG, respectively). Spirales
spore types that are represented by 40 and 35%
for NAG and AG respectively. Retinaculiaperti
was less observed (4 and 8%) (Figure 5). The
present study is the first report of the diversity and
ecological characterization of streptomycetes from
agricultural and non-agricultural soils and provides
new data on the populations of Streptomyces
influenced by soil chemical properties as well as
contributes to a better understanding of the dyna-
mic of soil microbial communities. Further studies
are needed to elucidate the biocontrol activity of
isolates and their role in agricultural fields with dif-
 Oskay 1003

Figure 3. Similarities (in percentage) of Streptomyces diversity between AG and NAG soil
samples.

Figure 4. Color series (in percentage) of Streptomyces isolated from AG and

NAG soil samples.
1004 Sci. Res. Essays
Figure 5. Spore bearing hyphae types (in percentage) of Streptomyces
isolated from AG and NAG soil samples (Representative of total 50
different streptomycete isolates were analyzed using cover clip method
with ISP2 and ISP4).
ferent crop species.
REFERENCES
Anderson AS, Wellington MHE (2001). The taxonomy of Streptomyces
and related genera. Int. J. Syst. Evol. Microbiol. 51: 797-814.
Conn KL, Leci E (1998). A Quantitative Method for Determining Soil
Populations of Streptomyces and Differentiating Potential Potato
Scab–Inducing Strains. Plant Dis. 82: 631-638.
Cross T (1989). Growth and Examination of Actinomycetes Some
Guidelines. In Bergey's Manual of Systematic Bacteriology. Williams
and Wilkins Company, Baltimore 4: 2340-2343.
Dolotkeldieva T, Totubaeva N (2006). Biodiversity of Streptomyces of
high-mountainous ecosystems of Kyrgyzstan and its biotechnological
potential. Anton. van Leeuwen. 89: 325-328.
El-Nakeeb MA, Lechevalier HA (1963). Selective Isolation of Aerobic
Actinomycetes Appl. Microbiol. 11: 75-77.
Grishko VN, Syshchikova OV (2009). Streptomyces Communities in
Soils Polluted with Heavy Metals. Eurasian Soil Sci. 42: 217-224.
Katsifas EA, Giannoutsou EP, Karagouni AD (1999). Diversity of
streptomycetes among specific Greek terrestrial Ecosystems. Let.
Appl. Microbiol. 29: 48-51.
Kuster E, Williams ST (1964). Selection of media for isolation of
Streptomycetes. Nature 202: 928–929.
Lartey RT (2006). Dynamics of Soil Flora and Fauna in Biological
Control of Soil Inhabiting Plant Pathogens. Plant Pathol. J. 5: 125-
142.
Lechevalier MP, Lechevalier HA (1970). Chemical composition as a
criterion in the classification of aerobic actinomycetes. Int. J. Syst.
Bacteriol. 20: 435–443.
Li HX, Li QR, Jiang CL (1996). Diversity of Soil Actinomycetes in
Yunnan, China. Appl. Environ. Microbiol. 62: 244-248.
Locci R (1989). Streptomyces and related Genera. In: Bergey’s Manual
of Systematic Bacteriology, Williams and Wilkins Company, Baltimore
4: 2451-2508.
Martinez Blanco E, Little CR, Davelos Baines AL (2007). Variation in
antibiotic inhibitory abilities among streptomycetes from south Texas
agricultural soils. Soil Biol. Biochem. 39: 268-275.
Mitra A, Chandra Santra S, Mukherjee J (2008). Distribution of
actinomycetes, their antagonistic behaviour and the physico-chemical
characteristics of the world’s largest tidal mangrove forest. Appl.
Microbiol. Biotechnol. 80: 685-695.
Parfitt RL, Yeates GW, Ross DJ, Mackay AD, Budding PJ (2005). Rela-
tionships between soil biota, nitrogen and phosphorus availability and
pasture growth under organic and conventional management. Appl.
Soil Ecol. 28: 1-13.
Prauser H (1964). Aptness and application of color for exact description
of colors of Streptomyces. Z. Allg. Mikrobiol. 4: 95-98.
Ryan J, Garabet S, Harmsen K, Rashid A (1996). A Soil and Plant
Analysis Manual Adapted for the West Asia and North Africa Region.
Icarda, Aleppo, Syria. 140: 87-90.
Saadoun I, Gharaibeh R (2003). The Streptomyces flora of Badia region
of Jordan and its potential as a source of antibiotics active against
antibiotic-resistant bacteria. J. Arid Environ. 53: 365-371.
Scheffer F, Schachtschabel P (1966). Lehrbuch der Boden kunde
Ferdinand Enke Verlag: Stuttgard p. 473.
Schlichting E, Blume HP (1966). Bodenkundliches praktikum. Verlag
Paul Paney, Hamburg und Berlin. pp. 121-125.
Schloter M, Bach H.J, Metz S, Sehy U, Munch JC (2003). Influence of
precision farming on the microbial community structure and functions
in nitrogen turnover. Agric. Ecosys. Environ. 98: 295-304.
Shirling EB, Gottlieb D (1966). Methods for characterization of
Streptomyces species. Int. J. Syst. Bacteriol. 16: 313-340.
Thakur D, Yadav A, Gogoi BK, Bora TC (2007). Isolation and screening
of Streptomyces in soil of protected forest areas from the states of
Assam and Tripura, India, for antimicrobial metabolites. Mycmed. 17:
242-249.
Vargas Gil S, Meriles J, Conforto C, Figoni G, Basanta M, Lovera E,
March GJ (2009a). Field assessment of soil biological and chemical
quality in response to crop management practices. World J.
Microbiol. Biotechnol. 25: 439-448.
Vargas Gil S, Pastor S, March GJ (2009b). Quantitative isolation of
biocontrol agents Trichoderma spp., Gliocladium spp. and action-
mycetes from soil with culture media. Microbiol. Res. 164: 196-205.
Vieira FCS, Nahas E (2005). Comparison of microbial numbers in soils
by using various culture media and temperatures. Microbiol.
Res.160:197-202.
Vijayakumar R, Muthukumar C, Thajuddin N, Panneerselvam A,
Saravanamuthu R (2007). Studies on the diversity of actinomycetes
in the Palk Strait region of Bay of Bengal, India. Actinomycetologica
21: 59-65.
Waksman SA (1961). The Actinomycetes: Classification, identification
and descriptions of genera and species. The Williams and Wilkins
Company, Baltimore 2: 61-292.
Watve MG, Tickoo R, Maithili M, Jog B, Bhole D (2001). How many
antibiotics are produced by the genus Streptomyces? Arch. Microbiol.
176: 386-390.
 Oskay 1005

Williams ST, Goodfellow M, Alderson G (1989). Genus Streptomyces Zenova GM, Gryadunova AA, Pozdnyakov AI, Zvyagintsev DG (2008).
Waksman and Henrici 1943, 339AL. In: Williams ST, Sharpe ME, Holt Aerobic and Microaerophilic Actinomycetes of Typical Agropeat and
JG (ed), Bergey's Manual of Systematic Bacteriology, Williams and Peat Soils. Eurasian Soil Sci. 41: 210-214.
Wilkins, Baltimore 4: 2452-2492.
Williams ST, Goodfellow M, Alderson G, Wellington EMH, Sneath PHA,
Sackin MJ (1983). Numerical classification of Streptomyces and
related genera. J. General Microbiol. 129: 1743-1813.